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Bystrow’s Paradox – gills, fossils, and the fish-to-tetrapod transition


Rainer R. Schoch, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Germany. E-mail:


Schoch, R.R. and Witzmann, F. 2011. Bystrow’s Paradox – gills, fossils, and the fish-to-tetrapod transition. —Acta Zoologica (Stockholm) 92: 251–265.

The issue of which breathing mechanism was used by the earliest tetrapods is still unsolved. Recent discoveries of stem tetrapods suggest the presence of internal gills and fish-like underwater breathing. The same osteological features were used by Bystrow to infer a salamander-like breathing through external gills in temnospondyl amphibians. This apparent contradiction – here called Bystrow’s Paradox – is resolved by reviewing the primary fossil evidence and the anatomy of the two gill types in extant taxa. Rather unexpectedly, we find that internal gills were present in a range of early crown tetrapods (temnospondyls), based on the anatomy of gill lamellae and location of branchial arteries on the ventral side of gill arch elements (ceratobranchials). Although it remains to be clarified which components are homologous in external and internal gills, both gill types are likely to have been present in Palaeozoic tetrapods – internal gills in aquatic adults of some taxa, and external gills in the larvae of these taxa and in larvae of numerous forms with terrestrial adults, which resorbed the external gills after the larval phase. Future developmental studies will hopefully clarify which mechanistic pathways are involved in gill formation and how these might have evolved.